Weismann August
Studies in the Theory of Descent, Volume I
II. Seasonal Dimorphism and Climatic Variation
If, as I have attempted to show, seasonal dimorphism originates through the slow operation of a changed summer climate, then is this phenomenon nothing else than the splitting up of a species into two climatic varieties in the same district, and we may expect to find various connexions between ordinary simple climatic variation and seasonal dimorphism. Cases indeed occur in which seasonal dimorphism and climatic variation pass into each other, and are interwoven in such a manner that the insight into the origin and nature of seasonal dimorphism gained experimentally finds confirmation. Before I go more closely into this subject, however, it is necessary to come to an understanding as to the conception “climatic variation,” for this term is often very arbitrarily applied to quite dissimilar phenomena.
According to my view there should be a sharp distinction made between climatic and local varieties. The former should comprehend only such cases as originate through the direct action of climatic influences; while under the general designation of “local forms,” should be comprised all variations which have their origin in other causes – such, for example, as in the indirect action of the external conditions of life, or in circumstances which do not owe their present existence to climate and external conditions, but rather to those geological changes which produce isolation. Thus, for instance, ancient species elsewhere long extinct might be preserved in certain parts of the earth by the protecting influence of isolation, whilst others which immigrated in a state of variability might become transformed into local varieties in such regions through the action of ‘amixia,’24 i.e. by not being allowed to cross with their companion forms existing in the other portions of their habitat. In single cases it may be difficult, or for the present impossible, to decide whether we have before us a climatic form, or a local form arising from other causes; but for this very reason we should be cautious in defining climatic variation.
The statement that climatic forms, in the true sense of the word, do exist is well known to me, and has been made unhesitatingly by all zoologists; indeed, a number of authentically observed facts might be produced, which prove that quite constant changes in a species may be brought about by the direct action of changed climatic conditions. With butterflies it is in many cases possible to separate pure climatic varieties from other local forms, inasmuch as we are dealing with only unimportant changes and not with those of biological value, so that natural selection may at the outset be excluded as the cause of the changes in question. Then again the sharply defined geographical distribution climatically governed, often furnishes evidence of transition forms in districts lying between two climatic extremes.
In the following attempt to make clear the relationship between simple climatic variation and seasonal dimorphism, I shall concern myself only with such undoubted climatic varieties. A case of this kind, in which the winter form of a seasonally dimorphic butterfly occurs in other habitats as the only form, i.e., as a climatic variety, has already been adduced in a former paragraph. I allude to the case of Pieris Napi, the winter form of which seasonally dimorphic species occurs in the temperate plains of Europe, whilst in Lapland and the Alps it is commonly found as a monomorphic climatic variety which is a higher development of the winter type, viz., the var. Bryoniæ.
Very analogous is the case of Euchloe Belia, a butterfly likewise belonging to the Pierinæ, which extends from the Mediterranean countries to the middle of France, and everywhere manifests a very sharply pronounced seasonal dimorphism. Its summer form was, until quite recently, described as a distinct species, E. Ausonia. Staudinger was the first to prove by breeding that the supposed two species were genetically related.25 This species, in addition to being found in the countries named, occurs also at a little spot in the Alps in the neighbourhood of the Simplon Pass. Owing to the short summer of the Alpine climate the species has in this locality but one annual brood, which bears the characters of the winter form, modified in all cases by the coarser thickly scattered hairs of the body (peculiar to many Alpine butterflies,) and some other slight differences. The var. Simplonia is thus in the Alps a simple climatic variety, whilst in the plains of Spain and the South of France it appears as the winter form of a seasonally dimorphic species.
This Euchloe var. Simplonia obviously corresponds to the var. Bryoniæ of Pieris Napi, and it is highly probable that this form of E. Belia must likewise be regarded as the parent-form of the species surviving from the glacial epoch, although it cannot be asserted, as can be done in the case of Bryoniæ, that the type has undergone no change since that epoch, for Bryoniæ from Lapland is identical with the Alpine form,26 whilst E. Simplonia does not appear to occur in Polar countries.
Very interesting also is the case of Polyommatus Phlæas, Linn., one of our commonest Lycænidæ, which has a very wide distribution, extending from Lapland to Spain and Sicily.27 If we compare specimens of this beautiful copper-coloured butterfly from Lapland with those from Germany, no constant difference can be detected; the insect has, however, but one annual generation in Lapland, whilst in Germany it is double-brooded; but the winter and summer generations resemble each other completely, and specimens which had been caught in spring on the Ligurian coast were likewise similarly coloured to those from Sardinia. (Fig. 21, Plate II.). According to these facts we might believe this species to be extraordinarily indifferent to climatic influence; but the South European summer generation differs to a not inconsiderable extent from the winter generation just mentioned, the brilliant coppery lustre being nearly covered with a thick sprinkling of black scales. (Plate II., Fig. 22.) The species has thus become seasonally dimorphic under the influence of the warm southern climate, although this is not the case in Germany where it also has two generations in the year.28 No one who is acquainted only with the Sardinian summer form, and not with the winter form of that place, would hesitate to regard the former as a climatic variety of our P. Phlæas; or, conversely, the north German form as a climatic variety of the southern summer form – according as he accepts the one or the other as the primary form of the species.
Still more complex are the conditions in another species of Lycænidæ, Plebeius Agestis (= Alexis Scop.), which presents a double seasonal dimorphism. This butterfly appears in three forms; in Germany A and B alternate with each other as winter and summer forms, whilst in Italy B and C succeed each other as winter and summer forms. The form B thus occurs in both climates, appearing as the summer form in Germany and as the winter form in Italy. The German winter variety A, is entirely absent in Italy (as I know from numerous specimens which I have caught), whilst the Italian summer form, on the other hand, (var. Allous, Gerh.), does not occur in Germany. The distinctions between the three forms are sufficiently striking. The form A (Fig. 18, Plate II.) is blackish-brown on the upper side, and has in the most strongly marked specimens only a trace of narrow red spots round the borders; whilst the form B (Fig. 19, Plate II.) is ornamented with vivid red border spots; and C (Fig. 20, Plate II.) is distinguished from B by the strong yellowish-brown of the under side. If we had before us only the German winter and the Italian summer forms, we should, without doubt, regard them as climatic varieties; but they are connected by the form B, interpolated in the course of the development of both, and the two extremes thus maintain the character of mere seasonal forms.
III. Nature of the Causes producing Climatic Varieties
It has been shown that the phenomenon of seasonal dimorphism has the same proximate cause as climatic variation, viz. change of climate, and that it must be regarded as identical in nature with climatic variation, being distinguished from ordinary, or, as I have designated it, simple (monomorphic) climatic variation by the fact that, besides the new form produced by change of climate, the old form continues to exist in genetic connexion with it, so that old and new forms alternate with each other according to the season.
Two further questions now present themselves for investigation, viz. (1) by what means does change of climate induce a change in the marking and colouring of a butterfly? and (2) to what extent does the climatic action determine the nature of the change?
With regard to the former question, it must, in the first place, be decided whether the true effect of climatic change lies in the action of a high or low temperature on the organism, or whether it may not perhaps be produced by the accelerated development caused by a high temperature, and the retarded development caused by a low temperature. Other factors belonging to the category of external conditions of life which are included in the term “climate” may be disregarded, as they are of no importance in these cases. The question under consideration is difficult to decide, since, on the one hand, warmth and a short pupal period, and, on the other hand, cold and a long pupal period, are generally inseparably connected with each other; and without great caution one may easily be led into fallacies, by attributing to the influence of causes now acting that which is but the consequence of long inheritance.
When, in the case of Araschnia Levana, even in very cold summers, Prorsa, but never the Levana form, emerges, it would still be erroneous to conclude that it is only the shorter period of development of the winter generation, and not the summer warmth, which occasioned the formation of the Prorsa type. This new form of the species did not come suddenly into existence, but (as appears sufficiently from the foregoing experiments) originated in the course of many generations, during which summer warmth and a short development period were generally associated together. From the fact that the winter generation always produces Levana, even when the pupæ have not been exposed to cold but kept in a room, it would be equally erroneous to infer that the cold of winter had no influence in determining the type. In this case also the determining causes must have been in operation during innumerable generations. After the winter form of the species has become established throughout such a long period, it remains constant, even when the external influence which produced it (cold) is occasionally withdrawn.
Experiments cannot further assist us here, since we cannot observe throughout long periods of time; but there are certain observations, which to me appear decisive. When, both in Germany and Italy, we see Polyommatus Phlæas appearing in two generations, of which both the German ones are alike, whilst in Italy the summer brood is black, we cannot ascribe this fact to the influence of a shorter period of development, because this period is the same both in Germany and Italy (two annual generations), so that it can only be attributed to the higher temperature of summer.
Many similar cases might be adduced, but the one given suffices for proof. I am therefore of opinion that it is not the duration of the period of development which is the cause of change in the formation of climatic varieties of butterflies, but only the temperature to which the species is exposed during its pupal existence. In what manner, then, are we to conceive that warmth acts on the marking and colouring of a butterfly? This is a question which could only be completely answered by gaining an insight into the mysterious chemico-physiological processes by which the butterfly is formed in the chrysalis; and indeed only by such a complete insight into the most minute details, which are far beyond our scrutiny, could we arrive at, or even approximate to, an explanation of the development of any living organism. Nevertheless an important step can be taken towards the solution of this problem, by establishing that the change does not depend essentially upon the action of warmth, but upon the organism itself, as appears from the nature of the change in one and the same species.
If we compare the Italian summer form of Polyommatus Phlæas with its winter form, we shall find that the difference between them consists only in the brilliant coppery red colour of the latter being largely suffused in the summer form with black scales. When entomologists speak of a “black dusting” of the upper side of the wings, this statement must not of course be understood literally; the number of scales is the same in both forms, but in the summer variety they are mostly black, a comparatively small number being red. We might thus be inclined to infer that, owing to the high temperature, the chemistry of the material undergoing transformation in Phlæas is changed in such a manner that less red and more black pigment is produced. But the case is not so simple, as will appear evident when we consider the fact that the summer forms have not originated suddenly, but only in the course of numerous generations; and when we further compare the two seasonal forms in other species. Thus in Pieris Napi the winter is distinguished from the summer form, among other characters, by the strong black dusting of the base of the wings. But we cannot conclude from this that in the present case more black pigment is produced in the winter than in the summer form, for in the latter, although the base of the wings is white, their tips and the black spots on the fore-wings are larger and of a deeper black than in the winter form. The quantity of black pigment produced does not distinguish between the two forms, but the mode of its distribution upon the wings.
Even in the case of species the summer form of which really possesses far more black than the winter form, as, for instance, Araschnia Levana, one type cannot be derived from the other simply by the expansion of the black spots present, since on the same place where in Levana a black band crosses the wings, Prorsa, which otherwise possesses much more black, has a white line. (See Figs. 1–9, Plate I.) The intermediate forms which have been artificially produced by the action of cold on the summer generation present a graduated series, according as reversion is more or less complete; a black spot first appearing in the middle of the white band of Prorsa, and then becoming enlarged until, finally, in the perfect Levana it unites with another black triangle proceeding from the front of the band, and thus becomes fused into a black bar. The white band of Prorsa and the black band of Levana by no means correspond in position; in Prorsa quite a new pattern appears, which does not originate by a simple colour replacement of the Levana marking. In the present case, therefore, there is no doubt that the new form is not produced simply because a certain pigment (black) is formed in larger quantities, but because its mode of distribution is at the same time different, white appearing in some instances where black formerly existed, whilst in other cases the black remains. Whoever compares Prorsa with Levana will not fail to be struck with the remarkable change of marking produced by the direct action of external conditions.
The numerous intermediate forms which can be produced artificially appear to me to furnish a further proof of the gradual character of the transformation. Ancestral intermediate forms can only occur where they have once had a former existence in the phyletic series. Reversion may only take place completely in some particular characters, whilst in others the new form remains constant – this is in fact the ordinary form of reversion, and in this manner a mixture of characters might appear which never existed as a phyletic stage; but particular characters could certainly never appear unless they were normal to the species at some stage of phyletic development. Were this possible it would directly contradict the idea of reversion, according to which new characters never make their appearance, but only such as have already existed. If, therefore, the ancestral forms of A. Levana (which we designate as Porima) present a great number of transitional varieties, this leads to the conclusion that the species must have gone through a long series of stages of phyletic development before the summer generation had completely changed into Prorsa. The view of the slow cumulative action of climatic influences already submitted, is thus confirmed.
If warmth is thus without doubt the agency which has gradually changed the colour and marking of many of our butterflies, it sufficiently appears from what has just been said concerning the nature of the change that the chief part in the transmutation is not to be attributed to the agency in question, but to the organism which is affected by it. Induced by warmth, there begins a change in the ultimate processes of the matter undergoing transformation, which increases from generation to generation, and which not only consists in the appearance of the colouring matter in one place instead of another, but also in the replacement of yellow, in one place by white and in another by black, or in the transformation of black into white on some portions of the wings, whilst in others black remains. When we consider with what extreme fidelity the most insignificant details of marking are, in constant species of butterflies, transmitted from generation to generation, a total change of the kind under consideration cannot but appear surprising, and we should not explain it by the nature of the agency (warmth), but only by the nature of the species affected. The latter cannot react upon the warmth in the same manner that a solution of an iron salt reacts upon potassium ferrocyanide or upon sulphuretted hydrogen; the colouring matter of the butterfly’s wing which was previously black does not become blue or yellow, nor does that which was white become changed into black, but a new marking is developed from the existing one – or, as I may express it in more general terms, the species takes another course of development; the complicated chemico-physical processes in the matter composing the pupa become gradually modified in such a manner that, as the final result, a new marking and colouring of the butterfly is produced.
Further facts can be adduced in support of the view that in these processes it is the constitution of the species, and not the external agency (warmth), which plays the chief part. The latter, as Darwin has strikingly expressed it, rather performs the function of the spark which ignites a combustible substance, whilst the character of the combustion depends upon the nature of the explosive material. Were this not the case, increased warmth would always change a given colour29 in the same manner in all butterflies, and would therefore always give rise to the production of the same colour. But this does not occur; Polyommatus Phlæas, for example, becoming black in the south, whilst the red-brown Vanessa Urticæ becomes black in high northern latitudes, and many other cases well known to entomologists might be adduced.30 It indeed appears that species of similar physical constitution, i.e., nearly allied species, under similar climatic influences, change in an analogous manner. A beautiful example of this is furnished by our Pierinæ. Most of the species display seasonal dimorphism; as, for instance, Pieris Brassicæ, Rapæ, Napi, Krueperi, and Daplidice, Euchloe Belia and Belemia, and Leucophasia Sinapis, in all of which the difference between the winter and the summer forms is of a precisely similar nature. The former are characterized by a strong black dusting of the base of the wings, and by a blackish or green sprinkling of scales on the underside of the hind wings, while the latter have intensely black tips to the wings, and frequently also spots on the fore-wings.
Nothing can prove more strikingly, however, that in such cases everything depends upon the physical constitution, than the fact that in the same species the males become changed in a different manner to the females. The parent-form of Pieris Napi (var. Bryoniæ) offers an example. In all the Pierinæ secondary sexual differences are found, the males being differently marked to the females; the species are thus sexually dimorphic. Now the male of the Alpine and Polar var. Bryoniæ, which I conceive to be the ancestral form, is scarcely to be distinguished, as has already been mentioned, from the male of our German winter form (P. Napi, var. Vernalis), whilst the female differs considerably.31 The gradual climatic change which transformed the parent-form Bryoniæ into Napi has therefore exerted a much greater effect on the female than on the male. The external action on the two sexes was exactly the same, but the response of the organism was different, and the cause of the difference can only be sought for in the fine differences of physical constitution which distinguish the male from the female. If we are unable to define these differences precisely, we may nevertheless safely conclude from such observations that they exist.
I have given special prominence to this subject because, in my idea, Darwin ascribes too much power to sexual selection when he attributes the formation of secondary sexual characters to the sole action of this agency. The case of Bryoniæ teaches us that such characters may arise from purely innate causes; and until experiments have decided how far the influence of sexual selection extends, we are justified in believing that the sexual dimorphism of butterflies is due in great part to the differences of physical constitution between the sexes. It is quite different with such sexual characters as the stridulating organs of male Orthoptera which are of undoubted importance to that sex. These can certainly be attributed with great probability to sexual selection.
It may perhaps not be superfluous to adduce one more similar case, in which, however, the male and not the female is the most affected by climate. In our latitudes, as also in the extreme north, Polyommatus Phlæas, already so often mentioned, is perfectly similar in both sexes in colour and marking; and the same holds good for the winter generation of the south. The summer generation of the latter, however, exhibits a slight sexual dimorphism, the red of the fore wings of the female being less completely covered with black than in the male.
